Alternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov. (Pleosporales, Pleosporaceae): Two new species associated with leaf spot and blight diseases of date palm (Phoenix dactylifera L.)

«83 MycoKeys

MycoKeys 120: 295-315 (2025)
DOI: 10.3897/mycokeys.120.144245

Research Article

Alternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.
(Pleosporales, Pleosporaceae): Two new species associated with
leaf spot and blight diseases of date palm (Phoenix dactylifera L.)

Youssef Djellid'?©, Alla Eddine Mahamedi'®, Milan Spetik?®, Eliska Hakalova®®, Ales Eichmeier®®,
Micael Ferreira Mota Gongalves*®, Fouad Lamghari®®, Maryam Ali Saeed Mohamed Al Hmoudi®™®,

Akila Berraf-Tebbal'®

1 Laboratoire de Biologie des Systemes Microbiens (LBSM), Ecole Normale Supérieure Cheikh Mohamed EI Bachir El Ibrahimi de Kouba, 16308 Vieux-Kouba, Alger,

Algeria

an -F Ww ND

Département de Biologie, Faculté des Sciences de la Nature et de la Vie, et des Sciences de la Terre, Université de Ghardaia, 47000 Ghardaia, Algeria
Mendeleum - Institute of Genetics, Faculty of Horticulture, Mendel University in Brno, Valticka 334, 69144, Lednice, Czech Republic

CESAM, Departamento de Biologia, Universidade de Aveiro, 3810-193 Aveiro, Portugal

Fujairah Research Centre, Sakamkam Road, Fujairah 00000, United Arab Emirates

Corresponding author: Akila Berraf-Tebbal (berraf.a@hotmail. fr)

OPEN Qaceess

Academic editor: Thorsten Lumbsch
Received: 12 December 2024
Accepted: 2 July 2025

Published: 15 August 2025

Citation: Djellid Y, Mahamedi AE,
Spetik M, Hakalova E, Eichmeier

A, Goncalves MFM, Lamghari F, Al
Hmoudi MASM, Berraf-Tebbal A
(2025) Alternaria phoenicis sp. nov.
and Alternaria ouedrighensis sp.
nov. (Pleosporales, Pleosporaceae):
Two new species associated with
leaf spot and blight diseases of
date palm (Phoenix dactylifera L.).
Mycokeys 120: 295-315. https://doi.
org/10.3897/mycokeys.120.144245

Copyright: © Youssef Djellid et al.
This is an open access article distributed under
terms of the Creative Commons Attribution

License (Attribution 4.0 International - CC BY 4.0).

Abstract

Date palm (Phoenix dactylifera L.) is one of the oldest fruit crops grown in the semi-ar-
id and arid regions, playing significant ecological, environmental and socio-economic
roles. Recently, palm leaf spot and blight diseases have indeed emerged as significant
threats to phoeniciculture. They reduce yield and quality of dates leading to economic
losses. Therefore, a survey was conducted in four palm groves located in the Biskra and
Ghardaia provinces of Algeria. This investigation revealed two new Alternaria species
associated with leaf spot and blight symptoms on date palm. These newly identified
species are designated as A. phoenicis sp. nov. and A. ouedrighensis sp. nov., which
belong to the Ulocladioides and Embellisia sections, respectively. The isolates were phy-
logenetically identified using the key genetic markers of the genus including the large
subunit ribosomal DNA (LSU), internal transcribed spacer region of the ribosomal RNA
(ITS), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), RNA polymerase II subunit
(RPB2), translation elongation factor (TEF7) and plasma membrane (ATPase) genes and
illustrated based on the morphological characteristics.

Key words: Alternaria, leaf spot and blight diseases, Phoenix dactylifera L., phylogeny,
taxonomy

Introduction

The date palm (Phoenix dactylifera L.) is a dioecious perennial monocot in the
Arecaceae family, which comprises around 200 genera and 1500 species (Daw-
son 1982). It is a vital crop in desert regions, serving as a primary source of
food and trade from North Africa to India and across other subtropical areas
(Erskine et al. 2004). Notably, Algeria stands as the world’s third-largest date
producer, generating over 1.3 million tonnes annually, where date palms un-

295

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

derpin both traditional and modern Saharan agriculture (FAO 2023). However,
despite its economic significance, date palms are vulnerable to various patho-
genic fungi that can severely damage their stem, leaves, fruit, and root, leading
to substantial yield reductions (Bokhary 2010; El-Juhany 2010).

Among the fungi that impact date palms, A/ternaria emerges as a particu-
larly associated group with leaf spots and blight diseases in the Middle East
regions (El-Juhany 2010; Al-Sadi et al. 2012; Al-Nadabi et al. 2018). Alternaria,
a genus in the family Pleosporaceae, order Pleosporales, and phylum Asco-
mycota, was first described by Nees in 1816 with Alternaria tenuis designated
as the type species. Since then, the taxonomy of Alternaria has undertaken
significant revisions leading to the identification of numerous new species.
Presently, the genus comprises more than 360 species encompassing 29
sections (Simmons 2007; Woudenberg et al. 2013; Wijayawardene et al. 2020;
Li et al. 2023).

Species of A/ternaria occupy a wide range of ecological niches, occurring as
endophytes within apparently asymptomatic plant tissues, saprobes on various
substrates such as dead vegetation, paper, and food, and as pathogens that im-
pact both plants and animals, including humans, worldwide. This adaptability
enables them to thrive in diverse environments and interact with a wide range
of hosts (Blodgett et al. 2000; Larran et al. 2001; Feng et al. 2011; Qi et al. 2012;
Li et al. 2023; He et al. 2024).

The Alternaria genus consists of several phytopathogenic species that
cause diseases in a wide array of plants around the world, affecting key crops
such as cabbage, cauliflower, tomato, carrot, wheat, cucurbits and date palm
(Chaerani and Voorrips 2006; Logrieco et al. 2009; Rahimloo and Ghosta 2015;
Al-Nadabi et al. 2018; Jayawardena et al. 2019). These pathogens primarily
induce leaf spots and defoliation, characterized by necrotic lesions and yel-
lowing on leaves (Mac Kinon et al. 1999). They can also infect various plant
parts, including seedlings and fruits, leading to significant pre- and post-harvest
losses (Thomma 2003; Lawrence et al. 2016). Furthermore, Alternaria species
are recognized as seed-borne pathogens and are known for producing harmful
secondary metabolites, including phytotoxins and mycotoxins (Thomma 2003;
Simmons 2007; Gilardi et al. 2015; Lawrence et al. 2016; Chalkley 2020).

Alternaria genus includes morphologically diverse species traditionally iden-
tified by reproductive structures, sporulation patterns, and host interactions.
However, taxonomic classification has been debated due to species complex-
es and morphological variability influenced by environmental conditions and
host specificity (Elliot 1917; Fries 1832; Neergaard 1945; Joly 1964; Simmons
1967). Afterward, Simmons introduced practical criteria to standardize taxo-
nomic concepts for Alternaria species, focusing on colony and conidial mor-
phology (Simmons 2007). Therefore, in recent years, DNA sequencing of con-
served loci has massively improved the knowledge of fungal phylogeny. Several
studies have shown that phylogenetic analysis becomes a reliable approach
for species-level identification. The multilocus phylogeny using genetic regions
such as ITS, LSU, TEF7, RPB2, GAPDH and Alt-a1 combined with morphological
data are frequently used to resolve the taxonomy and identification of Alternar-
ia taxa. Thus, new species are increasingly described (Woudenberg et al. 2013;
Al Ghafri et al. 2019; Li et al. 2023; Aung et al. 2024; He et al. 2024; Jayawarde-
na et al. 2025).

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 296

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

During an investigation of Alternaria species in Algeria, two new taxa were
isolated from date palm (Phoenix dactylifera L.). This study used a polyphasic
approach, integrating both morphological and phylogenetic analyses, to char-
acterize these newly introduced taxa.

Materials and methods
Isolation and morphological studies

During 2017, a set of 40 samples comprising leaves, rachises, and leaflets with
spot lesions was collected from date palm trees in Ghardaia and Bechar prov-
inces, Algeria (Fig. 1). Plant material was carefully enclosed in paper bags and
transported to the laboratory. Subsequently, isolations were made from the
margin of symptomatic tissues. Small pieces (approx. 5 mm?) of rachis and
leaflets were surface sterilized in 5% sodium hypochlorite (NaOCl) for 8 and 4
min, respectively. They were rinsed thrice with sterile distilled water, then dried
with sterilized filter paper and placed onto the surface of potato dextrose agar
(PDA, Difco Laboratories). Plates were incubated at 25 °C until fungal growth
was perceived. The mycelium emerged from the fragments of the tissues were
transferred to new PDA plates and incubated under the same conditions.
Colony growth characteristics including surface and reverse appearance of the
culture were recorded after 7 days of incubation on 90 mm diameter PDA Petri
plates at 25 °C in darkness, following Li et al. (2022) and Luo et al. (2022). Growth
characteristics were determined on PDA plates incubated at different temperatures
from 5-40 °C at 5 °C intervals in the dark. Reference strains and specimens are
maintained at the Fungal Biodiversity Centre (CBS) and MEND-F fungal collections.
Fungal colonies were subcultured onto water agar medium, supplement-
ed with autoclaved poplar twigs to enhance sporulation (Santos and Phillips
2009). The cultures were maintained on a laboratory bench at approximate-
ly 20-25 °C, where they were exposed to diffused daylight. After two weeks,
observations of micromorphological features including conidial size, shape,
colour, striation, septation, conidiophores and conidiogenous cells mount-
ed into 100% lactic acid, were made using a Nikon Eclipse 80i microscope.
Photographs and measurements of fungal structures mounted in 100% lactic
acid were taken with a Nikon DSRi1 camera and the software NIS-Elements D
(Nikon). Thirty measurements per structure were performed and presented in
the quantitative format “(min—) low — up (~max) x (min-) low — up (~max) um
(av. Length mean + SD x Width mean + SD um)”, with full observed ranges (min-
imum-—maximum), typical ranges (low-up), and mean + standard deviation.

DNA extraction and sequencing

Genomic DNA of our isolates was extracted from 7-day-old mycelium grown on
PDA at 25 °C. The NucleoSpin Tissue kit (Macherey-Nagel, Diiren, Germany) was
used according to the manufacturer's instructions (https://www.mn-net.com).
Polymerase chain reaction amplifications of the large subunit ribosom-
al DNA (LSU), internal transcribed spacer of ribosomal DNA (ITS), parts of
glyceraldehyde-3-phosphate dehydrogenase (GAPDH), RNA polymerase II subunit
(RPB2), translation elongation factor (TEF7) and plasma membrane adenosine tri-

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 297

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

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Figure 1. Date palm tree with decline symptoms (a), rachis (b—-f) and leaflets (g, h) spots.

phosphatase (ATPase) genes were performed using primer pairs (Table 1). Poly-
merase chain reaction (PCR) mixtures and amplification conditions were conduct-
ed following the protocols described by Berbee et al. (1999) and Woudenberg et
al. (2013). PCR mixture contained 10 uM of primer, 200 UMdNTP 1xTaq reaction
buffer, 2 Units of AmpliTaq-DNA polymerase, 2.5 mMMgCl, and 10 ng of template
DNA for a final reaction volume of 25 ul. After amplification, the obtained PCR
amplicons were purified and sequenced by the company Eurofins (Germany).

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 098

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

Table 1. Primers used for PCR amplification and sequencing of Alternaria genes.

Genes Primers References
ITS ITS1 White et al. 1990
ITS4
DEPT EF1-728F Carbone and Kohn 1999
EF1-986R
RPB2 RPB2-5F2 Sung et al. 2007
RPB2-7cR Liu et al. 1999
GAPDH gpd 1 Berbee et al.1999
gpd 2
ATPase ATPDF1 Lawrence et al. 2013
ATPDR1
LSU LROR Rehner and Samuels 1994
LR7 Vilgalys and Hester 1990

Phylogenetic analysis

The obtained sequences of ITS, LSU, GAPDH, RPB2, TEF1 and ATPase regions
were checked and manually adjusted, when necessary, using BioEdit Sequence
Alignment Editor v.7.0.4.1 (Hall 1999). Sequence alignments were conduct-
ed through the online version of the multiple sequence alignment program
(MAFFT) v.7 (Katoh et al. 2019) using the default settings. Newly generated
sequences were deposited in GenBank (Table 2).

The phylogenetic analysis was conducted through Maximum Likelihood (ML)
and Maximum Parsimony (MP) methods using MEGA11 v.11.0.13 (Tamura et
al. 2021). The best-fit evolutionary model was determined automatically by
MEGA11 software. The ML analysis was conducted using heuristic searches
consisted of 1000 step utilizing the Nearest-Neighbour-Interchange (NNI) algo-
rithm with a Neighbour-Joining starting tree automatically generated. Whereas
for the MP analysis, the Tree-Bisection-Regrafting (TBR) algorithm was applied.
One thousand (1000) bootstrap replications were conducted to evaluate the
generated MP trees robustness. Cicatricea salina CBS 302.84 and Stemphylium
herbarum CBS 191.86 were used as outgroup taxa.

Results
Phylogenetic analyses

The PCR amplification of the LSU, ITS, GAPDH, RPB2, TEF7 and ATPase regions
yielded DNA fragments of about 1200, 600, 580, 950, 300 and 1200 bp, respec-
tively. Given the lack of the ATPase sequences for several species of the Alter-
naria genus and the majority of the species in the Ulocladioides sections, this
marker has been discarded from the phylogenetic analysis. Those, the con-
catenated LSU, ITS, GAPDH, RPB2, and TEF7 datasets consisted of 90 strains
corresponding to 78 species and two outgroup taxa. The alignment contained
2915 characters of which 2031 were constant, 23 were excluded, 161 were vari-
able and parsimony-uninformative and 700 were parsimony-informative. Max-
imum parsimony (MP) analyses of combined dataset produced a single most
parsimonious tree (score = 3577, Cl = 0.327, RI = 0.684 and HI = 0.673), which

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 299

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

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MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

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MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

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MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

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303

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

resulted in the identification of the strains. Furthermore, maximum likelihood
analyses on concatenated dataset yielded a phylogenetic tree (Fig. 2), which
was similar with maximum parsimony tree in terms of either major topology
or results. So, it was chosen for the phylogeny demonstration. Alignment and
phylogenetic trees were deposited at TreeBASE (ID: 31850).

In the phylogenetic analysis, all the clades corresponding to the Alternaria
sections were well resolved. Of these, 2 clades corresponding to the sections
Ulocladioides and Embellisia encompassed the strains of this study. The iso-
lates G11, A26 and A28 formed independent well-supported subclade with
high bootstrap support (100% ML and 94% MP; Fig. 2) within the section Ulo-
cladioides and were considered to represent a distinct species, which was de-
scribed here as Alternaria phoenicis sp. nov. The strain G92 clustered within
the section Embellisia with a high boostrap support (100% ML and 94% MP;
Fig. 1), but was phylogenetically different from the closest species within the
section. It represented a further distinct species, which was described here as
Alternaria ouedrighensis sp. nov. (Fig. 2).

Taxonomy

Alternaria phoenicis Y. Djellid, A. E. Mahamedi, F. Lamghari & A. Berraf-Tebbal,
sp. nov.

MycoBank No: 856854

Fig. 3

Type. ALGERIA + Ghardaia Province (32°10'18.174"N, 3°34'56.6976'E), on
symptomatic leaflet and rachis of Phoenix dactylifera L., 2017, Y Djellid,
(MEND-F-1166, holotype), ex-type culture CBS 152585.

Etymology. Named after the host genus (Phoenix) from which the fungus
was isolated.

Description. Colonies on PDA reaching 75 mm diam. after 7 d at 25 °C, circu-
lar, cottony with dense hyphae, off-white to light grey in the center, reverse buff
to dark brown in the center. Minimum temperature for growth 5 °C, optimum
25 °C, maximum 37 °C. On Potato dextrose agar (PDA; Fig. 3), conidiophores
arising directly from lateral of aerial hyphae, straight or curved, geniculate,
smooth-walled, with up to 5-septate, unbranched or with up to two branches,
pale brown; Conidia solitary, subcylindrical to obclavate, (18.1-) 21.4 - 29.1
(-38.8) x (7.4-) 9.7 — 12.8 (-14.8) um, (av. 25.3 + 3.9 x 11.2 + 1.6), non-beaked
with a narrow base, light brown, with some darkened middle transverse septa,
3-6 transverse septa, and 0-1 longitudinal or oblique septa per transverse seg-
ment; these primary conidia produce secondary conidiophores that consist ina
subapical extension from the conidial body. Sexual morph not observed.

Notes. Phylogenetically, this species grouped within Ulocladioides section
but was different from the closest species (A. malicola, A. preussii and A. cant-
lous) in a distinct lineage with 100% ML / 94% MP statistical support. Alternaria
phoenicis sp. nov. is different from its sister species A. malicola, A. preussii
and A. cantlous, based on sequences derived from five loci (Fig. 2). After con-
ducting a nucleotide pairwise comparison as recommended by Jeewon and
Hyde (2016), the present species can be distinguished from the closet spe-
cies A. malicola, A. preussii and A. cantlous. Based on GAPDH, RPB2 and TEF1

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 304

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

. consortialis CBS 198.67
. consortialis CBS 101229
. consortialis CBS 483.81
. consortialis CBS 121493
. atra CBS 102060
. atra CBS 195.67
A. microspora CBS 124391
A. zantedeschiae CBS 124113
A. castaneae CBS 124390
A. subcucurbitae CBS 123376
A. subcucurbitae CBS 121491
92/70 A. allii-tuberosi CBS 124112
-/81 a A. oblongo-obovoidea CBS 126317
A. cantlous MF P 262011
A. cantlous CBS 123007
67/57 A26
73/- Be aa CBS 152585 = G11 | Alternaria phoenicis sp. nov.
ra] °%*l nae
A. malicola CGMCC3.18704
A. preussii CBS 102062

sd > BB d

sect. Ulocladioides

84/81

100/99

sect. Pseudoulocladium 88/96

i A. chartarum CBS 200.67

A. chartarum CBS 115269

100/100 100/99 — A. eichhorniae CBS 489.92
96/99_61/80 A. betae-kenyensis CBS 118810
sect. Alternaria 100/00ta| 4: daucifolii CBS 118812

A. alternata CBS 916.96

94/93 A. salicicola MFLUCC 22-0072

LJ oe

A. vaccariicola CBS 118714

sect. Gypsophilae oi
A. juxtiseptata CBS 119673

sect. Helianthiinficientes A. vignae YZU 171715
) A. triangularis MAFF 246776

53/- A. photistica CBS 212.86
. Panax
eee A. panax CBS 482.81
e

A. thalictrigena CBS 121712

A. hyacinthi CBS 416.71
sect. Embellisioides 99/92 A. botryospora CBS 478.90
37/84 A. lolii CBS 115266

100/100

A. omanensis SQUCC 13580
100/100! A. omanensis SQUCC 15560

sect. Omanenses

Figure 2. Phylogenetic tree based on the maximum likelihood analysis of Alternaria species inferred from combined LSU,
ITS, GAPDH, RPB2 and TEF1. Maximum likelihood (ML) and maximum parcimony (MP) bootstrap values (2 50%) given at the
nodes (ML/MP) are computed at from 1000 replicates. The tree is rooted to Cicatricea salina (CBS 302.84) and Stemphyli-
um herbarum (CBS 191.86). The novel species are highlighted in bold. The monotypic lineages are indicated by black dots.

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 305

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

100/100

: A. capsici-annui CBS 504.74
sect. Ulocladium Pp

A. alternariae CBS 126989

e 98/61 A. argyranthemi CBS 116530

96/98 A. ershadii IRAN 3275C
ia A. parvicaespitosa LEP 014858
A. inflata FMR 16477
88/60 tt ie
A. brassicifolii CNU 111118

100/99

sect. Pseudoalternaria

100/99 , A. tellustris CBS 538.83
61/54] LL. A. chlamydosporigena CBS 341.71
ee A. embellisia CBS 339.71
1OCHOS A. radicicola NB830
CBS 152587 = G92 Alternaria ouedrighensis sp. nov.
A. soliaridae CBS 118387

sect. Embellisia
53/- l

[| 400/99 -— A. chlamydospora CBS 491.72
sect. Phragmosporae A. phragmospora CBS 274.70
100/100 A. limaciformis CBS 481.81

sect. Undifilum A. bornmuelleri DAOM 231361
e .A. dennisii CBS 476.90
100/100 Cicatricea salina CBS 302.84

Outgrou
Stemphylium herbarum CBS 191.86 ( GIOUP)

0.1

Figure 2. Continued.

genes, A. phoenicis sp. nov. has 7 bp differences (2%, no gap) in GAPDH, 1 bp
(1%, no gap) in RPB2 and 29 bp (7%, 6 gaps) in TEF7 when compared to A. mali-
cola. Alternaria preussii presents 5 bp differences (2%, no gap) in GAPDH and
11 bp (2%, no gap) in RPB2. However, A. cantlous shows 1 bp difference (1%,
no gap) in RPB2 and 29 bp (11%, 6 gaps) in TEF7. Morphologically, A. phoenicis
(Fig. 3) can be distinguished by having narrower conidia (7.4-14.8 um) com-
pared to the three closely related species: A. cantlous (7.4-14.8 um), A. preussii
(13.0-13.7 um), and A. malicola (8-16 um). In terms of length, its conidia are
shorter than those of A. cantlous (24-36 um) but longer when compared to
A. preussii (18.3-20.4 um). However, the conidial length of A. malicola (16-35
um) is comparable to that of A. phoenicis (18.1-38.8 um). Regarding the co-
nidial septation, A. phoenicis is characterized by multiple transverse septa (up
to 6). In contrast, its closely related species exhibit fewer transverse septa, up
to four in A. canlous and up to three in both A. preussii and A. malicola. Addi-
tionally, A. phoenicis has the fewest longitudinal septa (0-1), compared to A.
preussii (1-2), A. malicola (1-5), and A. canlous (0-2) (Runa et al. 2009; Wang
et al. 2010; Dang et al. 2018).

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 306

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

Figure 3. Morphology of Alternaria phoenicis. Colony on PDA after 7 days at 25 °C (A); Conidiophores and conidioge-
nouse cells (B, C); Conidia (D—M). Scale bars: 10 um.

Alternaria ouedrighensis, A. Berraf-Tebbal, A. E. Mahamedi, F. Lamghari, E.
Hakalova & Y. Djellid, sp. nov.

MycoBank No: 856855

Fig. 4

Type. ALGERIA * Biskra Province (34°44'16.0152"N, 5°22'10.1064"E), on symp-
tomatic leaf of Phoenix dactylifera L. 2017, Y Djellid (MEND-F-1168, holotype),
ex-type culture CBS 152587.

Etymology. Named after the valley of Oued Righ from which the fungus was
collected.

Description. Colonies on Potato dextrose agar (PDA) reaching 51 mm diam.
after 7 d at 25 °C, circular with concentric zonation of the growth, cottony with
dense hyphae, dark green, reverse dark brown, with a white halo at the edge. Min-
imum temperature for growth 5 °C, optimum 25 °C, maximum 37 °C. On PDA me-
dia (Fig. 4), conidiophores arising directly from lateral of aerial hyphae, straight or
curved, geniculate sympodial proliferation, verruculose thick-walled, with up to 12-
septate, unbranched or with up to three branches, light to dark brown; Conidia soli-
tary, ovoid to subcylindrical, (11.4-) 15.3 — 17.7 (-24.1) x (7.7—-) 9.9 — 10.9 (-12.9)

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 307

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

Figure 4. Morphology of A/ternaria ouedrighensis. Colony on PDA after 7 days at 25 °C (A); Conidiophores and conidiog-
enous cells (B, C); Conidia (D—M) Scale bars: 10 um.

um (av. 16.5 + 3.4 x 10.4 + 1.4), light brown to dark, rigid, and thickened transverse
septa, 1-3 transverse septa, and 0-1 longitudinal or oblique septa per transverse
segment; these primary conidia produce secondary conidiophores that consist of
a subapical extension from the conidial body. Sexual morph not observed.

Note. Phylogenetically A. ouedrighensis formed a sister branch with A.
embellisia, A. chlamydosporigena, A. radicicola and A. tellustris in Embellisia
section with 100% ML/100% MP bootstrap support. Alternaria ouedrighensis
sp. nov. is different from its sister species A. radicicola, A. embellisia and A.
tellustris based on sequences derived from five genes (Fig. 2). After conduct-
ing a nucleotide pairwise comparison as recommended by Jeewon and Hyde
(2016), the present species can be readily distinguished from the closet species

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 308

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

A. radicicola, A. embellisia and A. tellustris constructed on any of the LSU, ITS,
GAPDH, RPB2 and TEF17 genes, which has 3 bp difference (1%, no gap) inthe ITS
region, 6 bp (2%, no gap) in GAPDH, 16 pb (2%, no gap) in RPB2 and 15 bp (11%,
14 gap) in TEF7 when compared with A. radicicola, 1 bp (1%, no gap) in LSU, 6 bp
(2%, no gap) in ITS, 24 bp (4%, 1 gap) in GAPDH, 17 bp (2%, 1 gap) in RPB2, and
17 bp (11%, 13 gaps) in TEF7 when compared with A. embellisia, and 1 bp (1%,
no gap) in LSU, 3 bp (1%, no gap) in ITS, 12 bp (2%, 1 gap) in GAPDH, 17 bp (2%,
no gap) in RPB2 and 13 bp (9%, 14 gaps) in TEF7 with sister species A. tellustris.

Morphologically, A. ouedrighensis (Fig. 4) is distinct from the closest spe-
cies A. embellisia in conidial body size. Alternaria ouedrighensis has conidia
shorter and wider (11.4-24.1 x 7.7-12.9 um; av. 16.5 + 3.4 x 10.4 + 1.4 um)
than those of A. radicicola (20-38 x 7-10 um; Bessadat et al. 2025) and A.
embellisia (19.18-36.2 x 2.55-5.74 um; av. 12.64 x 4.34 um; Delgado Ortiz et
al. 2019). In addition, the conidia of A. ouedrighensis present fewer transverse
septa (1-3 transverse septa) than those of A. radicicola (3—5 transverse septa)
and A. embellisia (2 — 6 transverse septa). However, A. ouedrighensis presents
fewer longitudinal septa (0-1 septum) compared to A. embellisia (1 — 2 septa).

Discussion

In this study, two new species of Alternaria, A. phoenicis and A. ouedrighensis,
have been identified within the sections Ulocladioides and Embellisia, respec-
tively. These species were characterized and illustrated through comprehensive
morphological studies and a detailed polylocus phylogenetic analysis, which pro-
vides robust support for their classification within the genus. Both species are
associated with black spot and blight diseases symptoms on date palm (Phoenix
dactylifera L.). These diseases present a range of symptoms that can significant-
ly compromise the health and productivity of this host tree. Black spot disease
typically manifests as dark, circular lesions on the leaves, often surrounded by a
yellow halo, which may merge to form larger necrotic areas. This condition can
lead to premature fall of the leaves, thereby substantially reducing the photosyn-
thetic capacity of the plant (Elmer and Pscheidt 2014). While the blight disease
symptoms are characterized by rapid wilting and dieback of fronds. The affected
leaves exhibit browning that typically initiates at the tips and progresses inward,
leading to significant tissue necrosis and overall leaf decline, which can result
in wilting and dieback. These conditions can impact the structural integrity and
physiological function of the date palm (Namsi et al. 2019).

Alternaria phoenicis, the newly described species, forms a clearly separate
cluster within the section Ulocladioides, in the multi-locus phylogenetic trees
derived by analyses of a concatenated DNA sequence dataset. This section
encompasses a diverse group of species recognized for their significant eco-
logical roles and potential agricultural impacts. They are mostly known as sap-
rotrophs on a variety of host substrates as well as opportunistic human patho-
gens (Runa et al. 2009; Lawrence et al. 2016; Gannibal and Gomzhina 2024).
The Ulocladioides section was introduced in 2013 by Woudenberg et al. to ac-
commodate species previously classified under Ulocladium section. Thus, the
Ulocladioides section included 20 species typified by Alternaria cucurbitae. Re-
cently, Gannibal and Gomzhina (2024) assessed the species boundaries within
the Ulocladioides section by using multilocus phylogenetic analysis based on

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 309

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

the genealogical concordance phylogenetic species recognition (GCPSR) prin-
ciple. They also utilized the coalescent-based model Poisson tree processes
(PTP mPTP) and evaluated for the presence of recombination. As a result, they
suggested to eradicate nine species by joining four other species. Alternaria
atra and A. multiformis were united into the single species A. atra. Five species,
A. brassicae-pekinensis, A. consortialis, A. cucurbitae, A. obovoidea, and A. ter-
ricola, were combined in the species A. consortialis. Alternaria heterospora and
A. subcucurbitae were combined into one species, A. subcucurbitae. Alternaria
aspera, A. chartarum, A. concatenata, and A. septospora were combined into a
single species, A. chartarum. Morphologically, species within this section can
be identified by their short, geniculate conidiophores, with sympodial prolifera-
tions and obovoid, non-beaked conidia, with a narrow base, single or in chains
(Woudenberg et al. 2013; Li et al. 2023).

The second new species A. ouedrighensis is introduced and classified in
section Embellisia within the genus Alternaria. This section was established to
include previously described species under the genus Embellisia (Lawrence et
al. 2012). It is currently limited to only four species: A. embellisia Woudenb. &
Crous, the type species, along with A. chlamydosporigena Woudenb. & Crous,
A. tellustris (E.G. Simmons) Woudenb. & Crous and A. radicicola Bessadat &
Simoneau (Woudenberg et al. 2013; Li et al. 2023; Bessadat et al. 2025). Phy-
logenetic analyses revealed the close relationships among these four species
and highlight their evolutionary ties to other sections of the A/ternaria genus.
Notably, these species exhibit consistent morphological traits, including thick,
dark, and rigid conidial septa, along with a limited presence of longitudinal sep-
ta, which serve as identification keys. Additionally, members of this section
have been recognized as pathogens that impact various vegetable crops, par-
ticularly tomato and garlic (Simmons 2001; Woudenberg et al. 2013). Although
A. ouedrighensis is currently represented by a single isolate, its recognition as
a new taxon remains valid, consistent with previous studies (Crous et al. 2015;
Licking et al. 2021), that have formally described novel species based on dis-
tinct phylogenetic placement and unique morphological characteristics. Con-
sequently, it is necessary to set up larger surveys and isolations that include
more phoenicical production areas to better understand the diversity and intra-
specific variability within Alternaria species.

The identification of these new species not only enriches our understanding
of the diversity within the A/ternaria genus but also emphasizes the necessity
for effective management strategies to minimize the impact of this genus on
plant health and productivity.

Additional information
Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of Al

No use of Al was reported.

MycoKeys 120: 295-315 (2025), DOI: 10.3897/mycokeys.120.144245 310

Youssef Djellid et al.: A/ternaria phoenicis sp. nov. and Alternaria ouedrighensis sp. nov.

Funding

This study was supported by the Internal Grant of Mendel University in Brno with the
grant number IGA-ZF/2021-ST2003. Micael F.M. Gongalves thanks the FCT — Fundagao
para a Ciéncia e a Tecnologia I.P., under the project/grant UID/50006 + LA/P/0094/2020
(doi.org/10.54499/LA/P/0094/2020) and his contract 2022.00758.CEECIND/CP1720/
CT0051 (doi.org/10.54499/2022.00758.CEECIND/CP1720/CT0051). The authors
gratefully acknowledge the Fujairah Research Centre, UAE for the financial support.

Author contributions

Berraf-Tebbal A conceptualized and designed the study, Djellid Y, Mahamedi AE con-
ducted the investigation, Djellid Y, Gongalves MFM, Spetik M, Hakalova E, Al Hmoudi
MASM conducted the experiments, Mahamedi AE analysed the data, Berraf-Tebbal A,
Djellid Y, Mahamedi AE wrote and revised the original draft, Lamghari F, Eichmeier A
ensured the project administration, all authors reviewed the final manuscript.

Author ORCIDs

Youssef Djellid © https://orcid.org/0009-0007-6833-5439

Alla Eddine Mahamedi © https://orcid.org/0000-0002-9744-8973

Milan Spetik © https://orcid.org/0000-0001-7659-8852

Eliska Hakalova © https://orcid.org/0000-0002-5433-8993

Ales Eichmeier © https://orcid.org/0000-0001-7358-3903

Micael Ferreira Mota Goncalves © https://orcid.org/0000-0003-2295-3374

Fouad Lamghari © https://orcid.org/0009-0002-2789-2240

Maryam Ali Saeed Mohamed Al Hmoudi ® https://orcid.org/0009-0005-9207-4924
Akila Berraf-Tebbal © https://orcid.org/0000-0001-8517-8542

Data availability

All of the data that support the findings of this study are available in the main text.

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